Geometrical characteristics of eggs from 3 poultry species

L. C. Wang,*,1 Z. T. Ruan,*,1 Z. W. Wu,* Q. L. Yu,y F. Chen,y X. F. Zhang,* F. M. Zhang,z R. J. Linhardt,z

and Z. G. Liu*,2

*College of Animal Science and Technology, Zhejiang Agriculture & Forestry University, Lin’an 311300, Zhejiang,
China; yHangzhou Xiaoshan Chicken Breeding Co., Ltd., Hangzhou 311300, Zhejiang, China; and zDepartments of
Chemical and Biological Engineering, Chemistry and Chemical Biology, Biology, and Biomedical Engineering, Center
for Biotechnology and Interdisciplinary Studies, Rensselaer Polytechnic Institute, Troy 12180, New York, USA

ABSTRACT We studied the correlations between egg
geometrical parameters (i.e., egg shape index, sphericity,
geometric mean diameter, surface area, and volume) and
eggshell qualities, or the organic matrix in eggshell. Eggs
were collected from 5 poultry breeds belonging to 3 spe-
cies (commercial Hy-line Brown Chicken, Shaoxing
Duck, Jinding Duck, Taihu Goose, and Zhedong White
Goose). The geometrical parameters showed high vari-
ation among 3 species of poultry, and even between
breeds in the same species. The five geometrical param-
eters were grouped into 2 sets, one contained shape index
and sphericity, the other comprised geometric mean
diameter, surface area, and volume. The parameters in
the same set can be perfectly fitted to one another. Egg
weight, shell membrane weight, and calcified shell weight
were significantly correlated with geometric mean
diameter, surface area, and volume. In accordance with
Key words: egg, geometrical parameter, eggshell quality, eggshell matrix
false discovery rate–adjusted P value, both shell mem-
brane relative weight and calcified shell thickness showed
no significant correlations with any of the geometrical
parameters. However, the correlations between geomet-
rical parameters and other shell variables (calcified shell
weight, shell relative weight, calcified shell thickness
uniformity, and eggshell breaking strength) depend on
breed. Both constitutive proportions and percentage
contents of 3 eggshell matrix components (acid-insol-
uble, water-insoluble, and both acid and water
facultative-soluble matrix) had no effects on egg shape
and size. The correlations between the amounts of
various shell matrix, egg shape and size depend on breed
or species. This study provides a methodology and the
correlation between geometrical parameters and eggshell
qualities, and between geometrical parameters and
organic matrix components in calcified shells.
2021 Poultry Science 100:100965
https://doi.org/10.1016/j.psj.2020.12.062

INTRODUCTION parameters, such as egg volume, surface area, radius of

The calcified eggs of birds exhibit a variety of sizes and
shapes (Stoddard et al., 2017). It is a challenging task to
describe egg profiles accurately because the high vari-
ability of eggs exists not only among bird species, but
also among the individuals of the same flock. Precise
quantification of egg profiles can provide a powerful
tool for relevant biological studies on population, ecolog-
ical morphology, egg incubation, and development
(Barta and Szekely, 1997; Hutchinson, 2000; Stoddard
et al., 2017). The knowledge of egg contours is also mean-
ingful for the poultry industry. The egg geometrical
Ó 2020 The Authors. Published by Elsevier Inc. on behalf of Poultry
Science Association Inc. This is an open access article under the CC BY-
NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).
Received October 27, 2020.
Accepted December 21, 2020.
1
These authors contributed equally as co-first authors.
2
Corresponding author: liuzg007@zafu.edu.cn
curvature, etc., have been used in research on the numer-
ical simulation of the eggshell behavior under mechani-
cal loading (Perianu et al., 2010) or under thermal
treatments (Sabliov et al., 2002; Denys et al., 2003).
These geometrical parameters have also been used to
predict egg qualities, breeding egg hatchability, and
hatchling size (Severa et al., 2013).
Egg shape index (SI) is the classic geometrical param-
eter to describe eggshell shape (Sarica and Erensayin,
2004). In addition to SI, some other egg geometrical pa-
rameters and their mathematical equations have also
been derived to promote the description of egg contours
(Narushin, 1997). The surface area (S) and the geomet-
ric mean diameter of eggs (Dg) were given by Mohsenin
(Mohsenin, 1970); based on these results, the degree of
sphericity (F) and the volume (V) of eggs were further
deduced (Severa et al., 2013).
In terms of above 5 geometrical parameters, except
egg SI, other parameters are mainly used in the research

1
2 WANG ET AL.
of ecological morphology (Barta and Szekely, 1997;
Hutchinson, 2000; Stoddard et al., 2017), and in the nu-
merical simulation of the eggshell behavior in food pro-
cessing (Sabliov et al., 2002; Denys et al., 2003;
Perianu et al., 2010; Severa et al., 2013; Kumbar et al.,
2016), while seldom used in research of eggshell qualities.
The present study used eggs from chicken, duck, and
goose to probe the correlations between geometrical pa-
rameters and eggshell qualities, and between geometrical
parameters and organic matrix components in calcified
shells.
MATERIALS AND METHODS
Egg Samples Collection
Eggs were collected from 5 poultry breeds of 3 species
from Zhejiang Province, China (Table 1). Chicken eggs
were from 280-day-old commercial Hy-line Brown
Chicken cage-reared in Lin’an County. Duck eggs were
from 2 breeds, the first batch was 400-day-old purebred
Jinding Duck free-ranged in Yuhang County; the second
batch was 450-day-old purebred Shaoxing Duck cage-
reared in Zhuji County. The goose eggs were from 2
flocks: 300-day-old purebred Taihu Goose free-ranged
in Nanxun County; and 320-day-old purebred, free-
range Zhedong White Goose in Xiangshan County.
Eggs with normal shape and color were sampled on the
laying day from each flock in the same month. The sam-
ple size of each breed was showed in Table 1.
Measurement of Egg or Eggshell Variables
Egg Weight, Length, and Width The individual egg
weight was measured using balance; then egg length
and width were measured using caliper.
Eggshell Broken Strength The eggshell broken
strength was measured by eggshell strength gauge from
Fujihara Co. (Tokyo, Japan). Each chicken and duck
egg were vertically placed as sharp end head down and
blunt end head up so that the blunt end bore the pres-
sure. Goose eggs were too long to vertically place under
the sensor rod, so each egg was horizontally placed to
test the shell annulus with the maximum width bore
the pressure.
Eggshell Membrane Weight and Calcified Shell
Weight After eggshell breaking strength testing, each
egg was broken into halves, the shells were washed
with water, then rinsed with 5% EDTA for 30 min,
40 min, and 45 min. The cuticle covering the outer sur-
face of calcified shell was removed using a toothbrush,
then the shell membrane was carefully and manually
removed from calcified shell and collected. Finally, the
shell membranes and calcified shells were dried at 40 C
and weighed.
Calcified Shell Thickness and Shell Thickness
Integrity Using a modified method from the study by
Sun et al., 2012, each calcified shell was roughly marked
in 5 parts with equal length along the longitudinal axe,
that is blunt end, blunt zone, equator zone, sharp zone,
and sharp end. Six shell pieces were sampled around the
circumference of each part. The thickness of each shell
piece was measured with a digital micrometer. The
average thickness of total 30 shell pieces was calculated as
thickness of the eggshell. The eggshell thickness unifor-
mity was defined as the reciprocal of the coefficient of
variation (1/CV) of the thickness of 30 shell pieces of each
egg. The higher eggshell thickness uniformity means more
homogeneous of global eggshell thickness.
Extraction of Organic Matrix Components
From Calcified Shells
Eggs from 3 breeds, Hy-line Brown Chicken, Shaoxing
Duck, and Zhedong White Goose, were used for this
analysis. Based on the method (Liu et al., 2017), the
organic matrix components were extracted from each
calcified eggshell. Briefly, the eggshell was individually
powdered using mortar and pestle, then decalcified by
stirring with 10% acetic acid at 20 C for about 18 h.
The amount of 10% acetic acid used was about
22–25 mL per gram shell powder, and the acetic acid
was added stepwise as the proportion of 35:35:30, the
first 2 steps of decalcification were individually carried
out over 3–4 h, and the last time decalcification was
performed overnight.
After decalcification, the suspension was centrifuged
(fixed-angle rotor) at 23,500 ! g for 18 min and the pel-
let was washed twice with distilled water and freeze-
dried and it was designated as acid-insoluble matrix
(expressed as Matrix1). The supernatant (referred as
acid-soluble matrix) was repeatedly dialyzed four times
against 40 volumes of distilled water at 20 C using a
Spectra/Por 6 dialysis tubing bag (molecular weight cut-
off 8 kDa; Spectrum Labs, Rancho Dominguez, CA).
The sample was then centrifuged at 3,500 ! g for
40 min to obtain water-insoluble matrix (expressed as
Matrix2) and facultative-soluble matrix (both acid and
water soluble in supernatant, expressed as Matrix3).
The water-insoluble matrix (Matrix2) was freeze-dried,
and the facultative-soluble matrix (Matrix3) was
concentrated using a Millipore spin column (molecular
weight cutoff 8 kDa) and freeze-dried. The mass of
each dried matrix component was individually weighed.
Calculation of Egg Geometrical Parameters
Based on the measurements of egg length (L) and
width (W), 5 geometrical parameters were calculated:
The egg SI was calculated from the formula (Sarica
and Erensayin, 2004) SI 5 (W/L)*100. The geometric
mean diameter of eggs (Dg) was calculated from the for-
mula (Mohsenin, 1970) Dg 5 (L*W2)1/3
The surface area of eggs (S) was calculated from the
formula (Mohsenin, 1970; Baryeh and Mangope, 2003)
S 5 p * Dg2.
In accordance with the references (Severa et al., 2013;
Kumbar et al., 2016), the degree of sphericity of eggs (F)
5 (Dg/L) * 100; and the volume of eggs (V) 5 (p/6) *L
* W2.
 POULTRY EGG GEOMETRICAL SHAPE 3
Table 1. Five poultry breeds providing egg samples.
Poultry breeds Age (day old) Egg sample size Breed characterization Breeding site
Hy-line Chicken 280 100 commercial Lin’an County, Zhejiang Province, China
Jinding Duck 400 71 purebred Yuhang County, Zhejiang Province, China
Shaoxing Duck 450 97 purebred Zhuji County, Zhejiang Province, China
Taihu Goose 300 71 purebred Nanxun County, Zhejiang Province, China
Zhedong White Goose 320 84 purebred Xiangshan County, Zhejiang Province, China

Statistical Analysis
The software of one-way ANOVA in SPSS 19.0 was
used to analyze the difference of geometrical parameters
among eggs from various flocks. The software of Bivar-
iate Correlations in SPSS 19.0 was used to determine
the strength and direction of Pearson correlation coeffi-
cients between pairs of variables, including correlations
between geometrical parameters and eggshell or shell
matrix variables. The classical false discovery rate–
adjusted P-values were calculated using Benjamini &
Hochberg methods to perform multiple hypothesis test
correction (Benjamini and Hochberg, 1995). Statistical
values were expressed as mean 6 SD, and the threshold
of significant difference chosen for all analyses was set as
P , 0.05.
RESULTS
Geometrical Parameters of Eggs From the
Five Flocks
In the present study, parameters describing egg
geometrical, eggshell, and shell matrix characteriza-
tions were systematically determined (Table 2,
Appendix 1 and 2). And the analysis of egg geomet-
rical parameters (Table 2) showed: i) the differences
of egg SI among the 3 poultry species were very signif-
icant (P , 0.01); ii) the difference between 2 duck
breeds was also very significant (P , 0.01); iii) there
was no significant difference between 2 goose breeds
(P . 0.05). Egg SI is generally used as the geometrical
parameter to describe eggshell shape. Eggs can be
characterized as sharp, normal (standard), and round
if they have an SI value of ,72, between 72 and 76,
and .76, respectively (Sarica and Erensayin, 2004).
The SI of chicken eggs was greater than 76, eggs of
both duck breeds were between 72 and 76, and eggs
of both goose breeds were less than 72 (Table 2).
This suggests that the egg shapes among these 3 spe-
cies are highly variable with chicken eggs being
comparatively round, goose eggs are sharp, and duck
eggs have a standard shape. In addition, the sphericity
(F) of chicken eggs was significantly larger than eggs
from both duck breeds, and duck eggs were signifi-
cantly larger than the eggs of both goose breeds
(P , 0.01) (Table 2). This also shows that chicken
eggs are rounder, and goose eggs are sharper.
In addition, the other 3 geometrical parameters, that
is, geometric mean diameter, surface area, and volume
of eggs, also showed high variation among the 3 species
of poultry, and even between breeds of the same species.
These 3 parameters all show that goose eggs are signifi-
cantly larger than duck eggs (P , 0.01), and duck eggs
are significantly larger than chicken eggs (P , 0.01)
(Table 2). This is consistent with the weights of goose
eggs being much larger than duck eggs, and duck eggs
being larger than chicken eggs. Obviously, eggs of larger
weight would also have larger contours of diameter, sur-
face area, and volume.
Correlations Among the Five Geometrical
Parameters
The results of eggs from all 5 breeds showed that the
SI positively and significantly correlated with sphericity
(adjusted P , 0.01) (Table 3–5). The geometric mean
diameter, surface area, and volume of eggs, positively
and significantly correlated with each other (adjusted
P , 0.01) (Tables 2–4). Furthermore, the Pearson
correlation coefficients (r) of the above parameter
pairs were all equal to or close to 1 (Tables 2–4). The
squares of correlation coefficients (r2) were equal/close
to 1, suggesting the parameter pairs with high
correlation can be perfectly fitted by each other.
By contrast, only the SI or sphericity of Shaoxing
Duck eggs significantly and negatively correlated with
the geometric mean diameter, surface area, or the vol-
ume of eggs (adjusted P , 0.01) (Table 4). In the case
of eggs from the other 4 breeds, there were no significant
correlations between the SI or sphericity and the other 3
parameters (P . 0.05) (Tables 3–5).

Table 2. Geometrical characteristics of eggs from the 5 flocks.

Parameters Hy-line Chicken Jinding Duck Shaoxing Duck Taihu Goose Zhedong White Goose
Shape index (%) 77.19 6 2.35 A 73.26 6 2.83 C 75.12 6 2.19 B 67.75 6 2.88 D 68.00 6 2.22 D
Sphericity (%) 84.14 6 1.71 A 81.26 6 2.10 C 82.63 6 1.61 B 77.13 6 2.20 D 77.32 6 1.68 D
Geometric diameter (mm) 48.37 6 0.94 A 50.93 6 1.20 C 50.04 6 1.23 D 58.39 6 1.80 B 62.77 6 1.75 A
Surface area (cm2) 73.52 6 2.86 E 81.52 6 3.83 C 78.70 6 3.87 D 107.22 6 6.58 B 123.89 6 6.90 A
Volume (cm3) 59.31 6 3.47 E 69.27 6 4.88 C 65.71 6 4.86 D 104.54 6 9.61 B 129.82 6 10.80 A

Values in the same raw with same capital letters mean the difference between them is not significant (P . 0.05), and the different
capital letters mean the difference is very significant (P , 0.01).
4 WANG ET AL.

Table 3. Correlations among the 5 geometrical parameters of eggs from Hy-line Brown Chicken.
Geometrical parameters Statistics Shape index Sphericity Geometric mean diameter Surface area Volume
Shape index r 1 1.000 20.196 20.197 20.197
P 0.000** 0.059 0.059 0.058
Adjusted P 0.000** 0.060 0.060 0.060
Sphericity r 1 20.196 20.196 20.197
P 0.060 0.059 0.059
Adjusted P 0.060 0.060 0.060
Geometric mean diameter r 1 1.000 1.000
P 0.000** 0.000**
Adjusted P 0.000** 0.000**
Surface area r 1 1.000
P 0.000**
Adjusted P 0.000**
Volume r 1
P
Adjusted P

**Represent the correlations between parameters were very significant (P , 0.01).

Based the above analysis, it seems the 5 geometrical
parameters could be grouped into 2 correlated sets: 1)
SI and sphericity; 2) geometric mean diameter, surface
area, and volume, because the parameters in the same
set can be perfectly fitted with each other.
Correlations Between the Geometrical
Parameters and Eggshell Qualities
In terms of egg weight, shell membrane weight, and
calcified shell weight, the results on the 5 breeds all posi-
tively and significantly correlated with egg geometric
mean diameter, surface area, and volume (adjusted
P , 0.01) (Table 6). Membrane weight showed no signif-
icant correlation with egg SI and sphericity in all 5
breeds (P . 0.05) (Table 6). Both shell membrane rela-
tive weight (the membrane weight normalized by egg
weight) and eggshell thickness showed no significant cor-
relation with any geometrical parameters (adjusted
P . 0.05) (Table 6).
However, the correlations between egg weight or calci-
fied shell weight and egg SI or sphericity depended on
breed, that is, egg weight and calcified shell weight nega-
tively and significantly correlated with SI and sphericity
only in the case of Hy-line Brown Chicken and Shaoxing
Duck (adjusted P , 0.01) (Table 6). Furthermore, the
correlation between other shell variables and geomet-
rical parameters also depends on breed. Only Hy-line
Brown Chicken and Shaoxing Duck showed negative
and significant correlation between shell relative weight
(the calcified shell weight normalized by egg weight)
with geometric mean diameter, surface area, and volume
(adjusted P , 0.01) (Table 6), but no significant corre-
lation with egg SI and sphericity (P . 0.05) (Table 6).
In other 3 breeds, shell relative weight showed no signif-
icant correlation with all 5 geometrical parameters
(P . 0.05) (Table 6). Calcified shell thickness uniformity
of Hy-line Brown Chicken showed significant correlation
with all 5 geometrical parameters (Table 6). By contrast,
both duck breeds, shell thickness uniformity only posi-
tively and significantly correlated with SI and sphericity
(adjusted P , 0.01) (Table 6). In both goose breeds,
shell thickness uniformity showed no significant correla-
tions with all 5 geometrical parameters (P . 0.05)
(Table 6). Eggshell breaking strength, a very important
shell trait, showed no significant correlations with all 5
geometrical parameters in the case of Hy-line Brown
Chicken, both duck breeds, and Taihu Goose (adjusted
P . 0.05) (Table 6); however, it significantly and posi-
tively correlated with geometric mean diameter, surface

Table 4. Correlations among the 5 geometrical parameters of eggs from both duck breeds.
Geometrical parameters Statistics Shape index Sphericity Geometric mean diameter Surface area Volume
Shape index r 1 1.000 20.320 20.320 20.320
P 0.000** 0.002** 0.002** 0.002**
Adjusted P 0.000** 0.002** 0.002** 0.002**
Sphericity r 1.000 1 20.321 20.321 20.321
P 0.000** 0.002** 0.002** 0.002**
Adjusted P 0.000** 0.002** 0.002** 0.002**
Geometric mean diameter r 0.093 0.092 1 1.000 0.999
P 0.475 0.478 0.000** 0.000**
Adjusted P 0.504 0.504 0.000** 0.000**
Surface area r 0.091 0.090 1.000 1 1.000
P 0.488 0.491 0.000** 0.000**
Adjusted P 0.504 0.504 0.000** 0.000**
Volume r 0.088 0.087 0.999 1.000 1
P 0.500 0.504 0.000** 0.000**
Adjusted P 0.504 0.504 0.000** 0.000**

1) The upper triangular represents the results of Shaoxing Duck eggs, and the lower triangular represents the results of Jinding Duck
eggs; 2) ** represents the correlations between parameters were very significant (P , 0.01).
 POULTRY EGG GEOMETRICAL SHAPE 5
Table 5. Correlations among the 5 geometrical parameters of eggs from both goose breeds.
Geometrical parameters Statistics Shape index Sphericity Geometric mean diameter Surface area Volume
Shape index r 1 1.000 0.029 0.026 0.023
P 0.000** 0.809 0.829 0.848
Adjusted P 0.000** 0.854 0.854 0.854
Sphericity r 1.000 1 0.028 0.025 0.022
P 0.000** 0.814 0.834 0.854
Adjusted P 0.000** 0.854 0.854 0.854
Geometric mean diameter r 20.183 20.182 1 1.000 0.999
P 0.145 0.147 0.000** 0.000**
Adjusted P 0.155 0.155 0.000** 0.000**
Surface area r 20.181 20.180 1.000 1 1.000
P 0.148 0.151 0.000** 0.000**
Adjusted P 0.155 0.155 0.000** 0.000**
Volume r 20.180 20.179 0.999 1.000 1
P 0.152 0.155 0.000** 0.000**
Adjusted P 0.155 0.155 0.000** 0.000**

1) The upper triangular represented the results of Zhedong White Goose eggs, and the lower triangular represented the results of
Taihu Goose eggs. 2) ** Represents the correlations between parameters were very significant (P , 0.01).

area, and volume in Zhedong White Goose (adjusted DISCUSSION

P , 0.05) (Table 6).
Correlations Between the Geometrical
Parameters and Eggshell Organic Matrix
The calcified shell is a predominant contributor to the
mechanical properties of eggshell, and comprised about
95% of CaCO3 calcite crystals and organic matrix.
Because this was the first time the correlations between
the geometrical parameters and eggshell organic matrix
were probed, we systemically studied the amounts of
various matrix: Matrix1, acid-insoluble matrix; Matrix2:
water-insoluble matrix; Matrix3: acid and water
facultative-soluble matrix; and total matrix: extracted
from individual calcified shell. The analysis also included
the percentage contents of various matrices in per gram
of shell, and the constitutive proportions of 3 matrix
fractions in the total matrix. We used 3 breeds (Hy-line
Brown Chicken, Shaoxing Duck, and Zhedong White
Goose) for these analyses.
The results of all 3 breeds showed that the constitutive
proportions of 3 matrix fractions, or the percentage con-
tents of the 3 matrix fractions had no significant correla-
tions with the 5 geometrical parameters (P . 0.05)
(Table 7). Only in Hy-line Brown Chicken, the percent-
age content of total matrix was positively and signifi-
cantly correlated with geometric mean diameter,
surface area, and volume (adjusted P , 0.05)
(Table 7). These results suggest that both constitutive
proportions and percentage contents of the 3 matrix
fractions had no effect on egg shape and size. In Hy-
line Brown Chicken and Zhedong White Goose, the
amounts of 3 matrix and total matrix in an individual
eggshell were all significantly and positively correlated
with 5 geometrical parameters (adjusted P , 0.05)
(Table 7). However, the amount of any matrix in duck
shell showed no significant correlation with the 5
geometrical parameters (P . 0.05) (Table 7). These
data indicate that the correlations between the amount
of matrix in an individual eggshell and egg shape and
size depend on breed or species.
The precise determination of egg geometrical parame-
ters is critical for the construction of the correct mathe-
matical model needed for egg processing, manipulation,
transport applications, and predictions in poultry pro-
duction (Severa et al., 2013). A number of researchers
deduced mathematical equations to express the contours
of individual eggs (Narushin, 1997). The mathematical
equations, of the current 5 egg geometrical parameters,
are all adequately defined by only 2 variables: maximum
length and width of an egg. Because the determination of
egg length and width is much easier than the comparison
of geometrical parameters, these equations would be
very helpful for the evaluation of egg shape and size.
It has been reported that there are very significant cor-
relations between chicken egg surface area and volume,
but there is no significant dependence of both parame-
ters on egg SI (Severa et al., 2013). Similarly, in the pre-
sent study, eggs from 5 breeds belonging to 3 poultry
species, all showed that the 5 geometrical parameters
could be grouped into 2 correlated sets, one contained
egg SI and sphericity, and the other contained geometric
mean diameter, surface area, and volume. The parame-
ters belonging to the same set can be perfectly fitted
by each other with very high degree of correlation. How-
ever, except for Shaoxing Duck eggs, eggs of other 4
breeds showed that parameters in one set showed no sig-
nificant correlations with parameters in another set
(Tables 2–4).
Based on broad taxonomic scales of 1,400 bird species,
it has been reported that avian egg shape correlated with
flight ability, and adaptation for flight may have been
critical drivers of egg-shape variation in birds
(Stoddard et al., 2017). It is known that the wild ances-
tors of chicken, duck, and goose are Gallus gallus, Anas
platyrhynchos or Anas poecilorhyncha, and Anser cyg-
noides or Anser anser, respectively. The phylogenetic
relationship between duck and goose is much closer. In
the present study, the egg geometrical parameters
showed large variations among species; however, the
egg geometrical parameters of both duck breeds were
Table 6. Correlations between egg geometrical parameters and shell qualities of 5 breeds.

6
Geometrical Membrane Eggshell Eggshell Shell thickness
Breeds parameters Statistics Egg weight Membrane weight relative weight Eggshell weight relative weight thickness uniformity Shell strength
Hy-line Brown Shape index r 20.218 20.099 20.049 20.249 20.083 20.082 0.317 0.046
Chicken P 0.036* 0.345 0.636 0.015* 0.426 0.433 0.002** 0.657
Adjusted P 0.058 0.442 0.685 0.027* 0.503 0.503 0.005** 0.685
Sphericity r 20.218 20.099 20.049 20.249 20.083 20.082 0.317 0.046
P 0.036* 0.344 0.636 0.015* 0.424 0.431 0.002** 0.661
Adjusted P 0.058 0.442 0.685 0.027* 0.503 0.503 0.005** 0.685
Geometric mean r 0.974 0.362 0.155 0.418 20.316 20.043 20.270 20.111
diameter P 0.000** 0.000** 0.137 0.000** 0.002** 0.685 0.010** 0.288
Adjusted P 0.000** 0.001** 0.201 0.000** 0.005** 0.685 0.020* 0.389
Surface area r 0.974 0.363 0.156 0.416 20.317 20.044 20.268 20.112
P 0.000** 0.000** 0.134 0.000** 0.002** 0.673 0.010** 0.286
Adjusted P 0.000** 0.001** 0.201 0.000** 0.005** 0.685 0.020* 0.389
Volume r 0.974 0.365 0.158 0.415 20.319 20.046 20.265 20.112
P 0.000** 0.000** 0.131 0.000** 0.002** 0.660 0.011* 0.285
Adjusted P 0.000** 0.001** 0.201 0.000** 0.005** 0.685 0.020* 0.389
Jinding Duck Shape index r 0.071 0.038 20.015 0.073 0.006 0.091 0.418 0.120
P 0.588 0.768 0.911 0.581 0.966 0.495 0.001** 0.360
Adjusted P 0.779 0.969 0.969 0.779 0.969 0.735 0.003** 0.621
Sphericity r 0.070 0.037 20.016 0.072 0.005 0.090 0.417 0.120
P 0.592 0.778 0.901 0.586 0.969 0.500 0.001** 0.360
Adjusted P 0.779 0.969 0.969 0.779 0.969 0.735 0.003** 0.621

WANG ET AL.
Geometric mean r 0.975 0.337 20.103 0.580 0.027 0.252 20.006 0.138
diameter P 0.000** 0.008** 0.433 0.000** 0.841 0.054 0.965 0.294
Adjusted P 0.000** 0.019* 0.698 0.000** 0.969 0.121 0.969 0.567
Surface area r 0.976 0.339 20.101 0.578 0.024 0.250 20.007 0.136
P 0.000** 0.007** 0.442 0.000** 0.858 0.057 0.959 0.300
Adjusted P 0.000** 0.019* 0.698 0.000** 0.969 0.121 0.969 0.567
Volume r 0.976 0.340 20.100 0.577 0.022 0.248 20.007 0.134
P 0.000** 0.007** 0.447 0.000** 0.870 0.058 0.956 0.306
Adjusted P 0.000** 0.019* 0.698 0.000** 0.969 0.121 0.969 0.567
Shaoxing Shape index r 20.327 20.188 20.031 20.262 20.023 20.101 0.306 20.050
Duck P 0.002** 0.076 0.771 0.013* 0.830 0.341 0.003** 0.638
Adjusted P 0.004** 0.136 0.840 0.025* 0.840 0.482 0.007** 0.808
Sphericity r 20.327 20.188 20.031 20.261 20.022 20.100 0.306 20.049
P 0.002** 0.076 0.770 0.013* 0.840 0.347 0.003** 0.646
Adjusted P 0.004** 0.136 0.840 0.025* 0.840 0.482 0.007** 0.808
Geometric mean r 0.985 0.341 20.143 0.429 20.336 20.029 20.145 20.026
diameter P 0.000** 0.001** 0.178 0.000** 0.001** 0.785 0.173 0.807
Adjusted P 0.000** 0.003** 0.268 0.000** 0.003** 0.840 0.268 0.840
Surface area r 0.985 0.340 20.144 0.429 20.335 20.027 20.143 20.025
P 0.000** 0.001** 0.174 0.000** 0.001** 0.802 0.177 0.812
Adjusted P 0.000** 0.003** 0.268 0.000** 0.003** 0.840 0.268 0.840
Volume r 0.986 0.339 20.145 0.430 20.333 20.024 20.142 20.025
P 0.000** 0.001** 0.173 0.000** 0.001** 0.822 0.182 0.818
Adjusted P 0.000** 0.003** 0.268 0.000** 0.003** 0.840 0.268 0.840
Taihu Goose Shape index r 20.195 20.188 20.083 20.155 20.067 20.050 0.175 0.260
P 0.120 0.133 0.513 0.220 0.598 0.694 0.163 0.036*
Adjusted P 0.235 0.245 0.605 0.350 0.673 0.705 0.272 0.087
Sphericity r 20.194 20.186 20.081 20.154 20.066 20.048 0.175 0.261
P 0.122 0.137 0.520 0.224 0.606 0.705 0.163 0.035*
Adjusted P 0.235 0.245 0.605 0.350 0.673 0.705 0.272 0.087
Geometric mean r 0.986 0.509 20.133 0.552 20.219 0.060 20.120 0.145
diameter P 0.000** 0.000** 0.292 0.000** 0.082 0.635 0.339 0.249
Adjusted P 0.000** 0.000** 0.384 0.000** 0.170 0.681 0.413 0.370
Surface area r 0.987 0.509 20.133 0.552 20.221 0.058 20.121 0.143
P 0.000** 0.000** 0.291 0.000** 0.079 0.645 0.337 0.254
Adjusted P 0.000** 0.000** 0.384 0.000** 0.170 0.681 0.413 0.370
Volume r 0.987 0.509 20.133 0.553 20.223 0.057 20.122 0.142
P 0.000** 0.000** 0.292 0.000** 0.076 0.654 0.334 0.259
Adjusted P 0.000** 0.000** 0.384 0.000** 0.170 0.681 0.413 0.370
Zhedong White Shape index r 0.018 0.073 0.098 20.070 20.115 20.067 0.219 0.079
Goose P 0.877 0.536 0.406 0.553 0.331 0.571 0.065 0.511
Adjusted P 0.883 0.680 0.606 0.680 0.573 0.680 0.122 0.680
Sphericity r 0.017 0.072 0.097 20.071 20.114 20.067 0.218 0.079

POULTRY EGG GEOMETRICAL SHAPE

P 0.883 0.543 0.412 0.551 0.333 0.571 0.066 0.510
Adjusted P 0.883 0.680 0.606 0.680 0.573 0.680 0.122 0.680
Geometric mean r 0.988 0.530 0.111 0.679 20.053 0.253 0.024 0.320
diameter P 0.000** 0.000** 0.345 0.000** 0.656 0.030* 0.844 0.006**
Adjusted P 0.000** 0.000** 0.573 0.000** 0.729 0.064 0.883 0.015*
Surface area r 0.988 0.528 0.109 0.677 -0.056 0.250 0.022 0.318
P 0.000** 0.000** 0.356 0.000** 0.636 0.032* 0.856 0.006**
Adjusted P 0.000** 0.000** 0.573 0.000** 0.723 0.066 0.883 0.015*
Volume r 0.988 0.526 0.106 0.675 -0.059 0.247 0.021 0.316
P 0.000** 0.000** 0.367 0.000** 0.617 0.034* 0.863 0.007**
Adjusted P 0.000** 0.000** 0.573 0.000** 0.717 0.068 0.883 0.016*

*Represents the correlations between parameters were significant (P , 0.05); **represents the correlations were very significant (P , 0.01).

7
 8
Table 7. Correlations between egg geometrical parameters and shell matrix of 3 poultry species.
Geometrical Matrix 1 Matrix 2 Matrix 3 Matrix 1 Matrix 2 Matrix 3 Total matrix Matrix 1 Matrix 2 Matrix 3 Total matrix
Breeds parameters Statistics constitute constitute constitute content content content content amount amount amount amount
Hy-line Shape index R 20.002 20.035 0.103 0.088 0.051 0.131 0.095 20.040 20.062 0.034 20.077
Brown P 0.987 0.762 0.369 0.443 0.660 0.252 0.411 0.731 0.591 0.769 0.506
Chicken Adjusted P 0.987 0.846 0.655 0.696 0.825 0.478 0.685 0.846 0.825 0.846 0.760
Sphericity R 20.002 20.035 0.103 0.089 0.052 0.132 0.096 20.039 20.062 0.035 20.076
P 0.987 0.761 0.368 0.440 0.653 0.249 0.404 0.737 0.594 0.764 0.511
Adjusted P 0.987 0.846 0.655 0.696 0.825 0.478 0.685 0.846 0.825 0.846 0.760
Geometric mean R 0.172 20.178 20.007 0.194 0.054 0.183 0.301 0.519 0.215 0.349 0.521
diameter P 0.133 0.119 0.954 0.089 0.639 0.109 0.008** 0.000** 0.060 0.002** 0.000**
Adjusted P 0.279 0.277 0.987 0.272 0.825 0.277 0.036* 0.000** 0.220 0.011* 0.000**
Surface area R 0.171 20.177 20.005 0.195 0.055 0.185 0.302 0.519 0.216 0.350 0.521
P 0.135 0.120 0.964 0.087 0.631 0.105 0.008** 0.000** 0.059 0.002** 0.000**
Adjusted P 0.279 0.277 0.987 0.272 0.825 0.277 0.036* 0.000** 0.220 0.011* 0.000**
Volume R 0.170 20.177 20.004 0.196 0.057 0.187 0.304 0.519 0.216 0.352 0.521
P 0.137 0.121 0.973 0.085 0.622 0.100 0.007** 0.000** 0.059 0.002** 0.000**
Adjusted P 0.279 0.277 0.987 0.272 0.825 0.277 0.036* 0.000** 0.220 0.011* 0.000**
Shaoxing Shape index r 0.101 20.040 20.178 0.136 0.094 20.123 0.127 20.002 20.019 20.162 20.016
Duck P 0.374 0.729 0.113 0.232 0.408 0.278 0.263 0.987 0.867 0.151 0.887
Adjusted P 0.628 0.872 0.628 0.628 0.646 0.628 0.628 0.991 0.950 0.628 0.950

WANG ET AL.
Sphericity r 0.101 20.040 20.178 0.136 0.094 20.122 0.127 20.001 20.019 20.161 20.016
P 0.371 0.724 0.114 0.231 0.411 0.281 0.263 0.991 0.866 0.153 0.891
Adjusted P 0.628 0.872 0.628 0.628 0.646 0.628 0.628 0.991 0.950 0.628 0.950
Geometric mean r 0.040 20.054 0.100 20.122 20.171 0.069 20.157 0.112 20.012 0.169 0.103
diameter P 0.727 0.636 0.377 0.283 0.132 0.543 0.168 0.327 0.915 0.133 0.366
Adjusted P 0.872 0.855 0.628 0.628 0.628 0.786 0.628 0.628 0.950 0.628 0.628
Surface area r 0.040 20.054 0.101 20.123 20.171 0.070 20.158 0.111 20.013 0.170 0.102
P 0.725 0.635 0.374 0.278 0.131 0.540 0.165 0.331 0.912 0.131 0.371
Adjusted P 0.872 0.855 0.628 0.628 0.628 0.786 0.628 0.628 0.950 0.628 0.628
Volume r 0.040 20.054 0.101 20.125 20.171 0.070 20.160 0.110 20.013 0.170 0.101
P 0.725 0.637 0.371 0.272 0.131 0.540 0.160 0.337 0.910 0.131 0.377
Adjusted P 0.872 0.855 0.628 0.628 0.628 0.786 0.628 0.628 0.950 0.628 0.628
Zhedong Shape index r 20.184 0.167 0.175 20.116 0.067 0.124 20.092 20.095 0.004 0.110 20.075
White P 0.117 0.155 0.135 0.325 0.570 0.298 0.433 0.423 0.972 0.352 0.527
Goose Adjusted P 0.322 0.322 0.322 0.483 0.668 0.468 0.554 0.554 0.974 0.499 0.644
Sphericity r 20.185 0.168 0.176 20.118 0.067 0.124 20.094 20.096 0.004 0.110 20.076
P 0.114 0.152 0.133 0.319 0.571 0.298 0.426 0.418 0.974 0.354 0.522
Adjusted P 0.322 0.322 0.322 0.483 0.668 0.468 0.554 0.554 0.974 0.499 0.644
Geometric mean r 20.178 0.166 0.138 20.016 0.219 0.160 0.028 0.356 0.560 0.467 0.416
diameter P 0.129 0.158 0.242 0.894 0.060 0.181 0.810 0.002** 0.000** 0.000** 0.000**
Adjusted P 0.322 0.322 0.403 0.928 0.252 0.355 0.918 0.011* 0.000** 0.000** 0.002**
Surface area r 20.181 0.169 0.140 20.022 0.216 0.157 0.022 0.352 0.557 0.466 0.412
P 0.123 0.151 0.234 0.854 0.064 0.187 0.851 0.002** 0.000** 0.000** 0.000**
Adjusted P 0.322 0.322 0.402 0.921 0.252 0.355 0.921 0.011* 0.000** 0.000** 0.002**
Volume r 20.184 0.171 0.142 20.027 0.213 0.155 0.017 0.348 0.554 0.464 0.408
P 0.118 0.145 0.228 0.818 0.069 0.194 0.889 0.003** 0.000** 0.000** 0.000**
Adjusted P 0.322 0.322 0.402 0.918 0.252 0.356 0.928 0.012* 0.000** 0.000** 0.002**

*Represents the correlations between parameters were significant (P , 0.05); ** represents the correlations were very significant (P , 0.01).
 POULTRY EGG GEOMETRICAL SHAPE
much close to that of chicken eggs (Table 2). Moreover,
based on the SI, our results showed that chicken eggs
were comparatively round, goose eggs were much
sharper, and duck eggs had an intermediate shape
(Table 2). It was reported that spherical eggs might be
advantageous because that spherical eggs have a mini-
mal surface-area-to-volume ratio, and require the least
amount of shell material for a given volume (Gosler
et al., 2005). Hy-line Brown chicken are characterized
by a high laying rate, and a laying interval between 2
consecutive eggs of about 22–26 h. Both Jinding Duck
and Shaoxing Duck lay about 300 eggs in one laying
year (Zhang et al., 2016), whereas Taihu Goose and Zhe-
dong White Goose lay 4 clutches of eggs in one laying
year, each clutch, respectively, contains 15 and 11
eggs, and the laying interval is about 44 h (Li et al.,
2010; Yao et al., 2018). Therefore, in the case of domes-
tication, the flight ability of poultry has been increas-
ingly lost, and the egg shape may be mainly affected
by laying rate, laying interval, or material demand of
eggshell formation.
Analysis showed that egg weight, shell membrane
weight, and calcified shell weight of all 5 breeds signifi-
cantly correlated with 3 geometrical parameters, geo-
metric mean diameter, surface area, and volume,
among which, the egg weight can be perfectly fitted by
the 3 egg geometrical parameters with very high correla-
tion (r . 0.97) (Table 6). This suggests that heavier eggs
had a larger contour of geometric mean diameter, surface
area, and volume. In addition, the significant correla-
tions between shell membrane weight or calcified shell
weight and above 3 geometrical parameters might be
indirectly affected by egg weight, because there was sig-
nificant dependence of both membrane and shell weights
on the egg weight (Appendix 3). Furthermore, it was
assumed that egg shape (spherical, ellipticity) was fixed
by the shell membranes (Stoddard et al., 2017), this
assumption might based on membrane material proper-
ties because in present study, eggs of 5 breeds all showed
that both egg SI and sphericity were independent on
membrane weight (Table 6).
The calcified shell thickness seems uniform in the same
egg, but the shell thickness is inhomogeneous to some
extent at different points. Therefore, shell thickness uni-
formity was introduced (Sun et al., 2012). It has been re-
ported that the thickness uniformity of chicken eggshells
was very significantly and positively correlated with
breaking strength (Sun et al., 2012). Our results revealed
the correlations between shell thickness uniformity and
egg geometrical parameters: in the case of Hy-line Brown
Chicken and both duck breeds, eggshell thickness unifor-
mity was very significantly and positively correlated
with SI and sphericity (Table 6). However, in both goose
breeds, shell thickness uniformity showed no significant
correlation with SI and sphericity (Table 6). The SI of
chicken eggs was greater than 76, eggs of both duck
breeds were between 72 and 76, and eggs of both goose
breeds were less than 72 (Table 2). It seemed that the
significant dependence of shell thickness uniformity on
the egg SI and/or sphericity might be applicable for
9
eggs with a SI value of .72, but not applicable for
eggs with a SI value of ,72; however, it is noteworthy
that we studied eggs from 180-, 500-, and 700-day old
of Shaoxing Duck flocks, the SI values were all above
72, but in the case of 700-day-old duck, the eggshell
thickness uniformity showed no significant correlation
with SI and sphericity (Luo et al., 2020). Therefore,
the significant correlations between shell thickness uni-
formity and SI or sphericity may depend on species, or
ages.
Based on sampling 8,000 eggs throughout hens’ entire
laying cycle, it had been reported that chicken eggshell
breaking strength significantly correlated with egg SI
(r 5 0.364, P , 0.01) (Sirri et al., 2018). However, the
present results showed that chicken eggshell breaking
strength showed no significant correlation with any of
the geometrical parameters (Table 6). The difference re-
sults might be attributed to the different sample sizes or
sampling methods. Furthermore, in the case of similar
sample size, only goose breeds showed significant correla-
tions between eggshell breaking strength and partial
geometrical parameters, suggesting that significant cor-
relations between poultry egg geometrical parameters
and eggshell breaking strength might depend on species
or breeds.
The calcified shell of avian egg, a composite bio-
ceramic formed by CaCO3 calcite crystals and a
pervading organic matrix, is a predominant contrib-
utor to the mechanical properties of the eggshell. The
organic matrix mainly comprises proteins and proteo-
glycans. In the present study, the organic matrix in
calcified eggshells were dissociated into acid-
insoluble, water-insoluble, and facultative-soluble
(both acid and water soluble) components. We had
previously reported the significant correlations be-
tween both egg weight and calcified eggshell weight
and the percentage contents of all 4 groups of matrix
(3 matrix components and total matrix) in per gram
of shell, and the significant correlations between both
calcified eggshell weight and shell thickness and the
amounts of all 4 groups of matrix in an individual
calcified shell (Liu et al., 2017). However, the present
results showed that the constituent ratios of 3 matrix
components in the whole matrix, and the percentage
contents of 3 matrix components in per gram of shell
showed no significant correlations with any geomet-
rical parameters, this meant both the constituent ra-
tios and percentage contents of 3 matrix components
showed no significant effects on egg contour. The sig-
nificant correlations between the amounts of 4 groups
of matrix in an individual calcified shell and egg size
depend on species, because in the case of duck eggs,
there was no significant correlation between the
amounts of 4 groups of matrix and egg size.
CONCLUSIONS
The egg geometrical parameters, adequately defined
by egg length and width, would be very helpful for the
evaluation of egg shape and size.
10 WANG ET AL.
The five geometrical parameters may be grouped into
2 correlated sets, and be of high variation among 3 spe-
cies of poultry.
The significant correlations between egg geometrical
parameters and some eggshell or organic matrix vari-
ables depend on species or breeds.
ACKNOWLEDGMENTS
The authors thank the Zhejiang Major Scientific and
Technological Project for Breeding New Agricultural
Breeds of Livestock and Poultry (2016C02054-12), Qian-
jiang Special Expert Project of Hangzhou City of China
(201929), Science and Technology Guidance Project of
Hangzhou City of China (201907), Zhejiang Provincial
Team Project for Animal Husbandry Technology
(2021-2022), and Zhejiang Provincial Key Laboratory
Project of Livestock and Poultry Ecological Health
Breeding (KLGEH012) for supporting this research.
DISCLOSURES
The authors of Geometrical characteristics of eggs
from 3 poultry species declared that they have no con-
flicts of interest to this work.
SUPPLEMENTARY DATA
Supplementary data associated with this article can
be found in the online version at https://doi.org/10.1
016/j.psj.2020.12.062.
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